1

Sandy Koi 1

Butterflies versus Bulldozers 2

Conservation Concerns for South Florida: Pine Rockland Denizens in Peril 3

9173 SW 72 Ave, M-5, Miami, FL 33156 4

sandykoi2009@gmail.com 5

1+(954) 449-5428 6

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2

Butterflies vs. Bulldozers 8

Conservation Concerns for South Florida: 9

Pine Rockland Denizens in Peril 10

© 2015 by S. Koi 11

Abstract 12

Urban/natural interface areas embody the classic conflicts between human desires and 13

endangered species, and these clashes are occurring more frequently worldwide. The playing 14

field becomes more complicated because development invites increased pesticide use and 15

decreased biodiversity; displaced wildlife is fragmented into disjointed and often isolated 16

populations. Fewer intact wild lands remain for wildlife to re-populate or for researchers to 17

relocate the exiled animals. Domestic gardens may provide habitat for some creatures, but more 18

often than not, urban wildlife becomes increasingly wedged between the concrete wall of a 19

shopping plaza and a super-highway. Here I describe several complex skirmishes that arose 20

between humans and invertebrate wildlife, and events that eradicated several self-established 21

colonies of the imperiled Atala Hairstreak butterfly in Southeast Florida, as well as two other 22

events that would have destroyed Atala colonies if were it not for the intervention of 23

conservation-minded individuals. In addition, the on-going threat of continued development in 24

the endangered Southeast Florida pine rocklands has the potential to devastate isolated colonies 25

of the United States federally endangered butterflies, Bartram’s Scrub Hairstreak and Florida 26

Leafwing, as well as one of the few remaining wild colonies of the Atala Hairstreak. 27

Keywords: Atala Butterfly, Bartram’s Scrub Hairstreak, Florida Leafwing, pine 28

rocklands, habitat alteration, defaunation, biodiversity loss 29

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Introduction: Global Defaunation & Habitat Skirmishes 30

Biodiversity loss is occurring at an unprecedented rate, aptly named the “defaunation” of 31

our ecosystems by Dirzo et al. (2014), and the sixth great extinction is here. Higgins described 32

the mass destruction of biomes and the biota inherent within it as “ecocide” and urged global 33

legislation to force the United Nations to include ecocide as one of the ‘crimes against humanity’ 34

(Higgins 2010). 35

Urban/natural interface areas exemplify the classic conflicts between human desires and 36

endangered species, and these clashes are occurring more frequently worldwide. Land alteration 37

in urban development invites increased pesticide use, is associated with decreased biodiversity, 38

and displaced wildlife is left in disjointed, often isolated, populations (Hubbuch 1991; Culbert 39

1994; Culbert 1995(2012); Myers et al. 2000; LaBonte et al. 2001; Rondeau 2001; Kershenbaum 40

et al. 2011; Dirzo et al. 2014). This is especially significant when the ecosystems supporting 41

endangered, threatened, vulnerable and imperiled life forms are themselves under threat 42

(Hubbuch 1991; Culbert 1994; Culbert 1995(2012); Myers et al. 2000; LaBonte et al. 2001; 43

Rondeau 2001; Kershenbaum et al. 2011; Dirzo et al. 2014), as is the case with Miami-Dade 44

County pine rockland ecosystems in Southeast Florida (Miami-Dade 2007). When these 45

geographical areas are also recognized as biodiversity hotspots globally (Myers et al. 2000), this 46

devastation is a conservation concern of even greater import. 47

The foundations of life on this planet are upheld by the vast abundance and diversity of 48

invertebrates, but are largely ignored for the ecological services they perform (Prather et al. 49

2012; Dirzo et al. 2014). More than half of monitored invertebrate insect populations recently 50

show a nearly fifty percent decline in abundance (Dirzo et al. 2014). Less than 400 of the 1800 51

butterfly species on the planet have been formally assessed by the International Union for 52

Conservation of Nature and Natural Resources (IUCN) and no up-to-date inventory of all 53

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butterfly species exists (Lewis 2010). Seventy-five percent of global fruit, vegetable, and grain 54

food crops are pollinated by insects (Gallai et al. 2009; Xerces 2011). Insects are enormously 55

responsible for soil and water nutrient recycling, detritus and waste disposal, food for other 56

species (including humans and other large mammals) as well as beneficial insect species control 57

of pest species (LaBonte et al. 2001; Losey and Vaughan 2006; Gallai et al. 2009; Xerces 2011; 58

Prather et al. 2012). 59

Many insect species are ephemeral, and/or diminutive, and are therefore difficult to 60

monitor, and challenging to find, and may live in isolated fragmented habitats (LaBonte et al. 61

2001; Taron and Reis 2015). Areas bordering urban localities are especially vulnerable; LaBonte 62

et al. (2001) point out that mosquito control spraying in recreational areas, controlled burns to 63

maintain forest ecology, road building for access to locations within the forest, the consequent 64

changes in soil chemistry from compaction and erosion, and herbicide use to control unwanted 65

invasive plant species are possible causes of the extirpation of the Polites mardon butterfly and a 66

flightless beetle, Agonum belleri. Negotiating the environmental impacts of pesticides used to 67

manage or eliminate invertebrate pests, while simultaneously balancing wildlife and human 68

safety, is challenging (Pimentel, 2009, 2011, 2013; Bargar 2012; Hoang et al. 2011; Xerces 69

2011; Pimentel et al. 2013). Some butterfly species disappear in an urban environment and others 70

are able to breed and even flourish, but most butterfly species are negatively impacted by urban 71

development (Hardy and Dennis 1999). 72

Climate change has also expanded or reduced potential range of many insects, but 73

anthropogenically altered landscapes negatively impact an animals’ ability to transverse 74

inhospitable matrices, obstructing possible new establishment sites (Gaston 2009; Thomas 2011; 75

Dirzo et al. 2014). Butterflies that inhabit montane regions may become extirpated as their 76

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ecosystems warm up (Forister and Shapiro 2003; Gaston et al. 2009), and in other cases, the 77

butterflies may expand their range (Koi and Daniels 2015). In addition, it is becoming 78

increasing difficult to find suitable locations for re-location or assisted translocation because of 79

increased habitat degradation. Thomas (2011) points out that we may be reaching a point where 80

restoration of species is no longer a viable alternative because of the anthropogenic alteration of 81

the remaining landscapes. 82

Butterflies and moths are one of the insect orders with the longest running monitor data 83

available and show consistent declines in numbers and abundance (Minno 2010, 2012; 84

Schweitzer et al. 2011; Dirzo et al. 2014; Taron and Reis 2015). Dirzo et al. (2014) indicated that 85

species richness in lepidoptera is 7.6 times higher on average in undisturbed habitat than in 86

disturbed sites. 87

Entomologists know that the ecosystem requirements for insect survival differs 88

dramatically between species, but have only recently recognized that management plans for 89

restoring degraded ecosystems, or establishing connectivity between patches, varies significantly 90

and is dependent on whether the target species is identified as “megafauna” or “microfauna” 91

(Longcore and Osborne 2015). Microfauna may need less actual space, but the micro-92

requirements of that space may mandate more specificity (Longcore and Osborne 2015). 93

Habitats, host plants, and imperiled insects in Southeast Florida 94

Pine rocklands are unique to Southeast Florida, Cuba and the Bahamas, containing 95

distinctive plants and animals adapted to a severe environment. The sharp limestone substrate 96

contains little soil; solution holes appear arbitrarily and the ecosystem is subject to stochastic 97

weather events, such as hurricanes, tropical storms and subsequent salt-water intrusion. It is a 98

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harsh environment and the biota is adapted to frequent and unpredictable natural and managed 99

fire; annual cycles of drought revolve around annual cycles of heavy rain. 100

Categorized as globally endangered by the Florida Natural Areas Inventory (Florida), 101

pine rocklands are also listed as “Environmentally Endangered Lands” in Miami-Dade County, 102

Southeast Florida (Snyder et al. 1990; Miami-Dade 2007; Florida 2010). The ecosystem is 103

dominated by slash pines (Pinus elliottii var. densa), an understory of dense saw palmetto 104

(Serenoa repens), cabbage palm (Sabal palmetto), and hundreds of rare tropical herbaceous taxa 105

endemic to the ecosystem. The habitat is maintained by fires, occurring in 3 to 7 year cycles, 106

from both managed fire and natural lightning strikes (Snyder et al. 1990; Miami-Dade 2007; 107

Florida 2010). The environment requires fire to suppress exotic plant invasion, reduce pine duff 108

and recycle organic material; it also prevents an otherwise natural progression into tropical 109

hardwood hammock. 110

The organic matter recycled by fire is especially important in the nutrient-poor limestone 111

substrate. In addition, some plants are fire-dependent, requiring fire in order to sprout, set seed or 112

grow. The flora is highly adapted to the environmental stresses and the pine rocklands contain a 113

rich biodiversity in flora and fauna (Miami-Dade 2007; Florida 2010). 114

Butterflies that have historically lived in pine rocklands include Bartram’s Scrub 115

Hairstreak (Strymon acis bartrami), the Florida Leafwing (Anaea troglodyta floridalis) and the 116

Atala Hairstreak (Eumaeus atala), all hostplant specialists. The host for Bartram’s and the 117

Leafwing is pineland croton (Croton linearis), which is seldom found outside the remaining 118

vestiges of pine rocklands in Southeast Florida. It is unlikely to be found in urban gardens and 119

the proximity of those gardens to pine rockland fragments would practically be a pre-requisite 120

for hosting a Bartram’s Scrub Hairstreak or a Florida Leafwing in a backyard garden. On 121

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September 11, 2014, the US Fish and Wildlife Service placed Bartram’s Scrub Hairstreak and 122

the Florida Leafwing on the official federally-recognized Endangered list (USFWS 2014). 123

North America’s only native cycad, Zamia integrifolia (=pumila=floridana) L. 124

(Zamiaceae: Cycadales), commonly called coontie, grows equally well in a backyard garden, an 125

urban plot, tropical and temperate hardwood hammocks, coastal shorelines or pine rocklands. 126

Once found historically from southern Georgia to the Florida Keys, the starch industries of the 127

last century depleted wild populations of coontie almost to the point of extirpation, as the roots 128

were harvested to make mildew-resistant flour, important in the humid sub-tropical environment 129

of Southeast Florida (Coile 2000; Coile and Gardner 2003). Coontie is still listed as 130

“Threatened” in wild natural areas (Coile and Gardner 2003; Donaldson 2013). 131

Cycads worldwide are currently perilously endangered or threatened (Oberpreiler 1995a, 132

1995b, 1995c, 2004; Chemnick et al. 2002; Donaldson 2003; Stevenson 2010); causes include 133

legal and illegal collection, removal of seed heads and root calyxes for food and/or “bush 134

medicines,” and the two biggest on-going threats: unsustainable trade and habitat loss 135

(Oberpreiler 1995a, 1995b, 1995c, 2004; Donaldson 2003). The Great Cycadian butterfly, 136

Eumaeus childrenae, G. Gray (1832) was recently declared endangered by a regional 137

conservation organization in Mexico, due to depletion of its six known cycad hostplants, all of 138

which are listed by IUCN as threatened, vulnerable or endangered (Donaldson 2003; Hernández-139

Baz and Rodríguez-Vargas 2014). Both legal and illegal trade impact the cycad colonies, many 140

of which are isolated and highly vulnerable, regardless of legislation and laws designed to 141

protect them (Oberpreiler 1995a, 1995b, 1995c, 2004; Chemnick et al. 2002; Donaldson 2003). 142

However, Florida’s native coontie has made a strong recovery in the landscaping industry 143

during the past twenty years, as nurseries and homeowners discovered how well-adapted the 144

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plant is to Southeast Florida’s diverse ecosystems and stochastic weather cycles (Haynes 2000; 145

Dehgan 2002; Culbert 1995(2010)). With increasing urban development and world nursery trade, 146

many more exotic non-native cycads have been added to the repertoire available to landscape 147

planners. Herbivory of the exotic cycads are a pest management concern of economic importance 148

as the plants are produced in highly specialized south Miami-Dade nurseries and are considered a 149

valuable addition to a prestigious landscape (i.e., the non-native cycads are not generally utilized 150

along highway median strips or shopping plaza parking lots). However, our native cycad has 151

shown itself to be hardy and resilient enough to be successfully utilized along mall parking lots 152

and highway medians. 153

Although it is generally accepted as beneficial that the Atala has increased its range and 154

distribution because of increased use of cycads for landscaping, the adult butterflies have 155

expanded their oviposition choices during the past twenty years as well. That expansion includes 156

many of the non-native introduced ornamental cycads found in south Florida’s botanical and 157

domestic gardens (Hubbuch 1991; Hammer 1985). The larvae are able to successfully complete 158

their life cycle on many, if not most, of these non-native plants (Hubbuch 1991; Hammer 1995; 159

Koi 2013, 2015). 160

This expansion into exotic Cycadales, many of which are extremely rare and valuable, 161

has in turn increased potential struggles between home-owners, botanists, property managers and 162

city planners with park managers, biologists, entomologists and conservationists as the Atala 163

butterfly larvae attack these urban cycads (Hubbuch 1991; Culbert 1994; Oberpreiler 1995a, 164

1995b, 1995c; Culbert 1995,2012). 165

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The reduction of the native cycads, as previously noted, caused the drastic decline in the 167

Atala butterfly’s population by the 1930’s, and by 1951 it was thought to be extinct (Klots 1951). 168

Unbeknownst to almost everyone, however, the butterfly had survived in isolated refugia (Koi 169

2013). Among the reasons that the Atala butterfly has made a dramatic recovery from near-170

extinction is the increased use of coontie, and other cycads in landscaping, as well as increased 171

conservation and restoration projects by biologists, botanists, park managers, scientists and 172

concerned citizens. 173

South Florida’s Butterflies Meet Anthropogenic Landscape Alterations 174

Unlike Bartram’s Scrub Hairstreak and the Florida Leafwing, whose hostplants are not 175

found in domestic gardens or landscaping practices, the Atala butterfly has utilized the increased 176

availability of native and non-native cycads found in ornamental landscapes. Currently, over 300 177

isolated and ephemeral populations of the butterfly are extant from Palm Beach to Miami-Dade 178

counties, most of which are harbored in private gardens rather than the few remaining natural 179

areas or parks (Koi 2004, 2013, 2015). The butterfly exhibits a classic crash-eruption cycle and 180

establishes ephemeral colonies in gardens and refugia wherever it finds suitable host and nectar 181

resources. These transient Atala colony sites may re-establish after being absent for as long as 8 182

years (Koi, unpublished). 183

The Atala butterfly has been documented in highly urbanized sites, as self-established, 184

relocated and introduced colonies (Koi 2004, 2013, 2015). One such colony self-established 185

along the shore of a heavily used urban coastal park in the City of Riviera Beach in Palm Beach 186

County, Florida; the site contained native and non-native cycads, as well as ‘weedy’ nectar 187

sources, such as Bidens alba L., and native palm trees. Neighborhood gardens surrounded the 188

colony as well. Fortunately the city zoning planner recognized the Atala butterfly and contacted 189

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the local butterfly club to let it be known that the city was going to raze the site for a complete 190

renovation. He asked for help to humanely remove the Atala colony and the host plants located 191

there until renovations were completed. 192

Volunteers quickly went into action and removed over sixty large mature coontie plants 193

from the site as well as over 50 adult Atala butterflies. The plants and butterflies, along with 194

immature life stages, were taken immediately to a mature butterfly garden that had been installed 195

years previously on a nearby college campus. Within minutes of being released in the garden, the 196

females were ovipositing on the newly moved coonties, which were planted later in the day. The 197

City of Riviera Beach has indicated that the Atala colony will be reinstated when the renovations 198

are completed. 199

Another self-established Atala colony was almost destroyed in a location in Broward 200

County, Florida a few months later. This was also an urban coastal site located in an abandoned 201

derelict parking lot, surrounded by heavily trafficked streets, businesses and residential housing. 202

This location contained more than a hundred very large healthy coontie plants which had been 203

installed between the parking spaces. There were large well-established trees that provided shade 204

and roosting spots, and abundant nectar was available in the alley-weeds, shrubs and trees along 205

the length of the neglected site.. 206

A volunteer had been monitoring this persistent Atala colony for nearly ten years and just 207

happened to be at the site counting the eggs, larvae and pupae of the Atala butterfly population 208

when she was approached and asked to explain what she was doing. After explaining the 209

butterfly monitoring method, the man revealed that he was the superintendent for a developer 210

that had recently bought the property and that they were about to raze it in a week. 211

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The superintendent was persuaded to allow volunteers three weeks to remove not only 212

the butterflies, but as many of the enormous coontie plants as possible, as well. The Atala 213

population erupted and crashed in normal ways over the years, but had persisted for many years. 214

Immediately, butterfly and native plant volunteers from Palm Beach, Broward and 215

Miami-Dade counties arrived during the three week respite period. Truckloads of coontie plants 216

were removed from the site; the rootstocks on these plants were deep and heavily entangled. It 217

was back-breaking work that took days to accomplish; the plants may have grown even larger 218

had they not been closed in by concrete berms. It is not known how long the parking lot had been 219

empty and unused, or how long ago the coontie was planted. 220

The adult butterflies were collected and released at a safe haven in Miami, but the 221

immatures (eggs, larvae and pupae) were removed with the plants to the many new locations to 222

start new colonies. Any remaining adult butterflies dispersed by themselves to establish new sites 223

in gardens nearby. Volunteers graciously counted the immature insects so that we could have a 224

final count of the last denizens of this colony (see Table 1). 225

Ultimately, this removal was beneficial to a number of new sites: in Delray Beach itself, in the 226

Town of Wellington, and several new colony introductions and re-introductions took place via 227

the Palm Beach County Department of Environmental Resources, including Delray Oaks Natural 228

Area, Pondhawk Natural Area, High Ridge Scrub Natural Area and Jupiter Ridge. ZooMiami 229

also benefited with rescued coontie plants, and later over 250 host plants for the Atala, Bartram’s 230

Scrub Hairstreak, the Florida Leafwing and Florida Duskywing were planted a few weeks later. 231

Continued Wildlife-Human Conservation Conflicts 232

Meanwhile, yet another development is currently being planned in Miami-Dade County 233

on a property known as the “Richmond Tract,” located beside the ZooMiami. This entire 234

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property is the largest remaining pine rockland habitat in Miami-Dade County, outside of 235

Everglades National Park, and represents most of the less than 2% which remains. The 236

Richmond Tract also contains 260 taxa of native plants (Miami 2007), many endangered 237

(Miami-Dade 2007; Florida 2010; USFWS 2014). The developer’s plans would bulldoze acres of 238

the remaining pine rocklands to build a housing development, a large chain retail store and 239

commercial shopping mall. 240

The Richmond Tract is currently home to not only the ‘imperiled’ Atala butterfly, but 241

also two other pineland denizens, Bartram’s Scrub Hairstreak and the Florida Leafwing, both 242

recently listed as Endangered by the United States Fish and Wildlife Service (USFWS 2014). In 243

addition, this property is adjacent to the Coastguard pinelands, where the Florida Duskywing 244

(Ephyriades brunnea floridensis), another of the 38 imperiled Southeast Florida butterfly 245

species, can be found. 246

There are also endangered Bonneted Bats (Eumops floridanus) on the site, and the 247

Richmond pine rocklands is one of the only places in the world where the miniscule green 248

iridescent Miami Tiger Beetle (Cicindela floridana) lives. Tiger beetles are named for their 249

aggressive stalking and lightning-fast predation and are highly sensitive to minor changes in their 250

environment (Pearson 2011). Tiger beetles occupy highly restricted habitats, and as such are 251

recognized as bioindicators (Pearson 2011). An emergency petition to USFWS has been filed to 252

request listing the Miami Tiger Beetle as federally endangered. 253

There are endangered plants located on the Richmond site, including the deltoid spurge 254

(Chamaesyce deltoidea), a rare prostrate plant which successfully stopped the destruction of the 255

nearby Rockdale pine rockland site in Miami many years ago. That site, a small triangular 256

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remnant of land, did not have the potential economic value that paving over the Richmond pine 257

rocklands could bring. 258

The Richmond tract contains Brickell-bush (Brickellia mosieri) and tiny polygala 259

(Polygala smallii), as well. Pine rocklands are a harsh environment in which to survive, but the 260

unique plants, insects and mammals have evolved over millennia together. Destroying them for 261

the sake of an amusement park and shopping plazas seems insane. 262

Butterflies, beetles and rare plants may be a hard sell against the possible economic 263

promise of a tourist-oriented theme parks and shopping plazas, but such unnecessarily 264

destructive development is surely a foolish direction for water-stressed and densely populated 265

Southeast Florida to travel. There are thousands of acres of derelict property in Miami-Dade 266

County that could be paved over for amusement parks without destroying the remaining remnant 267

natural areas. 268

And lastly, a thought from the world-renowned cycad botanist, Rolf Oberpreiler (1995c): 269

The natural insect fauna of cycads should not be brazenly dismissed as ‘pests.’ 270

These insects are a natural and mostly vital component of the environment of the 271

cycads, and their destruction can have severe impacts on the survival of the 272

plants. The most obvious examples in this regard are the pollinators, but other 273

insects may also play important roles in e.g., the disintegration of the cones and 274

release of the seeds, the decomposition of old cones, leaves and stems, the 275

recycling of nutrients, etc. . . .their survival is so inextricably attached to that of 276

the plants that extinction of their host plants will inevitably lead to the extinction 277

of these insect species. 278

It has been long recognized that destruction of the butterflies’ hostplants leads quickly to 279

either extinction, or extirpation of the butterflies (Koh et al. 2004; Gyllenberg and Hanski 1997). 280

“System-wide decline” begins with a loss of 30% of the natural vegetation cover (Fischer and 281

Lindenmayer 2007), and the cascading effect of species co-extinctions follows soon thereafter 282

(Koh et al. 2004). Especially for small, sedentary species such as beetles and butterflies, the 283

anthropogenic changes to the natural landscape, especially when coupled with climate change, 284

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can have a devastating effect on the animals (Walther et al. 2002; Koh et al. 2004; Fischer and 285

Lindenmayer 2007). As habitat loss becomes more pronounced, there are fewer suitable patches 286

from which re-colonization can occur, as well (Gyllenberg and Hanski 1997; Fischer and 287

Lindenmayer 2007). Habitat loss is considered the greatest threat to butterflies (Gyllenberg and 288

Hanski 1997; Koh et al. 2004; Fischer and Lindenmayer 2007; Forister et al. 2010; Thomas 289

2011). 290

Species at the highest risk for extinction are those with any or all of the following 291

characteristics: specialist hostplant use, limited dispersal ability (because of habitat alteration, 292

lack of mobility, body size, genetic behavior, or other factors), niche specialization, population 293

rarity or density, endemic to the locale, home range size, edge sensitivity, area requirements, 294

abiotic effects (humidity, temperature), complex community interactions (such as ants and 295

butterfly larvae), micro-habitat requirements, seasonal effects, connectivity of habitat patches, 296

and/or landscape structure needs (Gyllenberg and Hanski 1997; Koh et al. 2004; Fischer and 297

Lindenmayer 2007; Forister et al. 2010; Thomas 2011; Longcore and Osbourne 2015). 298

Because most extinctions occur in “island-like” environments with high concentrations of 299

endemic species (Fischer and Lindenmayer 2007; Thomas 2011), it is my hope that Miami-Dade 300

County, in the biogeographic “island-peninsula” of Southeast Florida, is not leading the way in 301

proving this further. 302

303

Acknowledgements: I thank journalist Al Sunshine for the title idea of this paper as we confront 304

the diminishing pine rocklands in Miami-Dade County, Florida. I acknowledge the ongoing 305

efforts of the volunteer citizen scientists who have helped with long-term data collection of E. 306

atala populations in Southeast Florida. I thank the University of Florida for permission to use 307

material from my thesis and my chair, Jaret Daniels. 308

309

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